Exotic grasses and nitrate enrichment alter soil carbon cycling along an urban–rural tropical forest gradient

Urban areas are expanding rapidly in tropical regions, with potential to alter ecosystem dynamics. In particular, exotic grasses and atmospheric nitrogen (N) deposition simultaneously affect tropical urbanized landscapes, with unknown effects on properties like soil carbon (C) storage. We hypothesized that (H1) soil nitrate (NO₃^?) is elevated nearer to the urban core, reflecting N deposition gradients. (H2) Exotic grasslands have elevated soil NO₃^? and decreased soil C relative to secondary forests, with higher N promoting decomposer activity. (H3) Exotic grasslands have greater seasonality in soil NO₃^? vs. secondary forests, due to higher sensitivity of grassland soil moisture to rainfall. We predicted that NO₃^? would be positively related to dissolved organic C (DOC) production via changes in decomposer activity. We measured six paired grassland/secondary forest sites along a tropical urban‐to‐rural gradient during the three dominant seasons (hurricane, dry, and early wet). We found that (1) soil NO₃^? was generally elevated nearer to the urban core, with particularly clear spatial trends for grasslands. (2) Exotic grasslands had lower soil C than secondary forests, which was related to elevated decomposer enzyme activities and soil respiration. Unexpectedly, soil NO₃^? was negatively related to enzyme activities, and was lower in grasslands than forests. (3) Grasslands had greater soil NO₃^? seasonality vs. forests, but this was not strongly linked to shifts in soil moisture or DOC. Our results suggest that exotic grasses in tropical regions are likely to drastically reduce soil C storage, but that N deposition may have an opposite effect via suppression of enzyme activities. However, soil NO₃^? accumulation here was higher in urban forests than grasslands, potentially related to of aboveground N interception. Net urban effects on C storage across tropical landscapes will likely vary depending on the mosaic of grass cover, rates of N deposition, and responses by local decomposer communities.     

Interactive Effects of Vegetation Type and Topographic Position on Nitrogen Availability and Loss in a Temperate Montane Ecosystem

Determining the fate of deposited nitrogen (N) in natural ecosystems remains a challenge. Heterogeneity of vegetation types and resulting plant–soil feedbacks interact with topo-hydrologic gradients to mediate spatial patterns of N availability and loss, yet net effects of variation in these two factors together across complex terrain remain unclear. Here we measured a suite of N-cycle pools and fluxes in sites that differed factorially in vegetation type (mixed forest vs. herbaceous) and topographic position (upslope vs. downslope) in a protected montane watershed near Salt Lake City, UT. Vegetation type was associated with large variation in N availability—herbaceous sites had larger NO₃ ^? pools, higher NO₃ ^?:NH₄ ⁺ ratios, higher nitrification potentials, lower soil C:N values, enriched δ¹⁵N values, and lower microbial biomass compared to forests, especially those upslope. Downslope sites tended to exhibit higher N availability and indicators of N-cycle openness, but patterns were moderated by vegetation type. In downslope forest, soil NO₃ ^? depth profiles and higher foliar N content suggested trees were accessing deep soil N and transferring it to the surface via litterfall, while more deep soil NO₃ ^? but no change in surface or foliar N suggested herbaceous cover was not N limited or deeper N pools were not accessible. Soil NO₃ ^? leaching from below the rooting zone closely tracked N availability, revealing a link between N status and hydrologic loss as well as an important role for roots in N retention. NO₃ ^? isotopes did not reveal a similar link for gaseous losses (that is, denitrification), instead reflecting nitrification and/or transport dynamics. Together, these results suggest a coupled ecological, topo-hydrologic perspective can help assess the fate of N in complex landscapes.     

Soil microbial community indices as predictors of soil solution chemistry and N leaching in Picea abies (L.) Karst. forests in S. Sweden

High rates of inorganic nitrogen (N) deposition or internal N turnover increases the risks of N loss from forests with negative effects on stream water quality. We hypothesized that soil fungi may be more important N sinks than bacteria, and thus examined the impact of soil microbial community composition on N leaching from forests. We studied 19 spruce stands to examine relationships between microbial community composition, stem growth, soil-, and lysimeter-collected soil solution characteristics, and N leaching. We used nitrate concentration in the soil solution below the rooting zone as an N leaching index and phospholipid fatty acid (PLFA) analysis for characterisation of microbial communities. Microbial community composition in the organic horizon and soil solution chemistry below the rooting zone was highly correlated. Stands with low concentrations of nitrate (NO₃ ^?) and aluminium (Al) had higher fungi: bacteria ratio compared with stands with higher concentrations of NO₃ ^? and Al. Stem growth and fungi: bacteria ratio explained 70 % of the variation in N and Al leaching. We identified three microbial predictors of variation in soil solution chemistry, of which the fungi: bacteria was the strongest. The other two were putative indicators of microbial C limitation, a condition known to stimulate N mineralisation and nitrification.     

Temperature sensitivity of soil carbon and nitrogen mineralization: impacts of nitrogen species and land use type

Background and aims

Climate warming, nitrogen (N) deposition and land use change are some of the drivers affecting ecosystem processes such as soil carbon (C) and N dynamics, yet the interactive effects of those drivers on ecosystem processes are poorly understood. This study aimed to understand mechanisms of interactive effects of temperature, form of N deposition and land use type on soil C and N mineralization.

Methods

We studied, in a laboratory incubation experiment, the effects of temperature (15 vs. 25 °C) and species of N deposition (NH₄ ⁺-N vs. NO₃ ^?-N) on soil CO₂ efflux, dissolved organic C (DOC) and N (DON), NH₄ ⁺-N, and NO₃ ^?-N concentrations using intact soil columns collected from adjacent forest and grassland ecosystems in north-central Alberta.

Results

Temperature and land use type interacted to affect soil CO₂ efflux, concentrations of DON, NH₄ ⁺-N and NO₃ ^?-N in most measurement times, with the higher incubation temperature resulted in the higher CO₂ efflux and NH₄ ⁺-N concentrations in forest soils and higher DON and NO₃ ^?-N concentrations in grassland soils. Temperature and land use type affected the cumulative soil CO₂ efflux, and DOC, DON, NH₄ ⁺-N and NO₃ ^?-N concentrations. The form of N added or its interaction with the other two factors did not affect any of the C and N cycling parameters.

Conclusions

Temperature and land use type were dominant factors affecting soil C loss, with the soil C in grassland soils more stable and resistant to temperature changes. The lack of short-term effects of the deposition of different N species on soil C and N mineralization suggest that maybe there was a threshold for the N effect to kick in and long-term experiments should be conducted to further elucidate the species of N deposition effects on soil C and N cycling in the studied systems.     

Stable N isotope composition of nitrate reflects N transformations during the passage of water through a montane rain forest in Ecuador

Knowledge of the fate of deposited N in the possibly N-limited, highly biodiverse north Andean forests is important because of the possible effects of N inputs on plant performance and species composition. We analyzed concentrations and fluxes of NO₃ ^??CN, NH₄ ⁺?CN and dissolved organic N (DON) in rainfall, throughfall, litter leachate, mineral soil solutions (0.15?C0.30 m depths) and stream water in a montane forest in Ecuador during four consecutive quarters and used the natural ¹⁵N abundance in NO₃ ^? during the passage of rain water through the ecosystem and bulk ??¹⁵N values in soil to detect N transformations. Depletion of ¹⁵N in NO₃ ^? and increased NO₃ ^??CN fluxes during the passage through the canopy and the organic layer indicated nitrification in these compartments. During leaching from the organic layer to mineral soil and stream, NO₃ ^? concentrations progressively decreased and were enriched in ¹⁵N but did not reach the ??¹⁵N values of solid phase organic matter (??¹⁵N = 5.6?C6.7??). This suggested a combination of nitrification and denitrification in mineral soil. In the wettest quarter, the ??¹⁵N value of NO₃ ^? in litter leachate was smaller (??¹⁵N = ?1.58??) than in the other quarters (??¹⁵N = ?9.38 ± SE 0.46??) probably because of reduced mineralization and associated fractionation against ¹⁵N. Nitrogen isotope fractionation of NO₃ ^? between litter leachate and stream water was smaller in the wettest period than in the other periods probably because of a higher rate of denitrification and continuous dilution by isotopically lighter NO₃ ^??CN from throughfall and nitrification in the organic layer during the wettest period. The stable N isotope composition of NO₃ ^? gave valuable indications of N transformations during the passage of water through the forest ecosystem from rainfall to the stream.     

Nitrogen deposition and forest nitrogen cycling along an urban–rural transect in southern China

There is increasing concern over the impact of atmospheric nitrogen (N) deposition on forest ecosystems in the tropical and subtropical areas. In this study, we quantified atmospheric N deposition and revealed current plant and soil N status in 14 forests along a 150 km urban to rural transect in southern China, with an emphasis on examining whether foliar δ¹⁵N can be used as an indicator of N saturation. Bulk deposition ranged from 16.2 to 38.2 kg N ha^?1 yr^?1, while the throughfall covered a larger range of 11.7–65.1 kg N ha^?1 yr^?1. Foliar N concentration, NO₃^? leaching to stream, and soil NO₃^? concentration were low and NO₃^? production was negligible in some rural forests, indicating that primary production in these forests may be limited by N supply. But all these N variables were enhanced in suburban and urban forests. Across the study transect, throughfall N input was correlated positively with soil nitrification and NO₃^? leaching to stream, and negatively with pH values in soil and stream water. Foliar δ¹⁵N was between ?6.6‰ and 0.7‰, and was negatively correlated with soil NO₃^? concentration and NO₃^? leaching to stream across the entire transect, demonstrating that an increased N supply does not necessarily increase forest δ¹⁵N values. We proposed several potential mechanism that could contribute to the δ¹⁵N pattern, including (1) increased plant uptake of ¹⁵N‐depleted soil NO₃^?, (2) foliage uptake of ¹⁵N‐depleted NH₄⁺, (3) increased utilization of soil inorganic N relative to dissolved organic N, and (4) increased fractionation during plant N uptake under higher soil N availability.     

Small differences in ombrotrophy control regional-scale variation in methane cycling among <Emphasis Type="Italic">Sphagnum</Emphasis>-dominated peatlands

Across northern Alberta, Canada, bogs experience periodic wildfire and, in the Fort McMurray region, are exposed to increasing atmospheric N deposition related to oil sands development. As the fire return interval shortens and/or growing season temperatures increase, the regional peatland CO₂–C sink across northern Alberta will likely decrease, but the magnitude of the decrease could be diminished if increasing atmospheric N deposition alters N cycling in a way that stimulates post-fire successional development in bogs. We quantified net ammonification, nitrification, and dissolved organic N (DON) production in surface peat along a post-fire chronosequence of five bogs where we also experimentally manipulated N deposition (no water controls plus 0, 10, and 20 kg N ha^?1 yr^?1 simulated deposition, as NH₄NO₃). Initial KCl-extractable NH₄⁺–N, NO₃^?–N and DON averaged 176?±?6, 54?±?0.2, and 3580?±?40 ng N cm^?3, respectively, with no consistent changes as a function of time since fire and no consistent effects of experimental N addition. Net ammonification, nitrification, and DON production averaged 3.8?±?0.3, 1.6?±?0.2, and 14.3?±?2.0 ng N cm^?3 d^?1, also with no consistent changes as a function of time since fire and no consistent effects of experimental N addition. Our hypothesis that N mineralization would be stimulated after fire because root death would create a pulse of labile soil organic C was not supported, most likely because ericaceous plant roots typically are not killed in boreal bog wildfires. The absence of any N mineralization response to experimental N addition is most likely a result of rapid immobilization of added NH₄⁺–N and NO₃^?–N in peat with a wide C:N ratio. In these boreal bogs, belowground N cycling is likely characterized by large DON pools that turn over relatively slowly and small DIN pools that turn over relatively rapidly. For Alberta bogs that have persisted at historically low N deposition values and begin to receive higher N deposition related to anthropogenic activities, peat N mineralization processes may be largely unaffected until the peat C:N ratio reaches a point that no longer favors immobilization of NH₄⁺–N and NO₃^?–N.     

Dissolved carbon and nitrogen losses from forests of the Oregon Cascades over a successional gradient

Ecologists have long used stream water chemistry records to infer hillslope processes, although a great deal of biogeochemical processing of soil water is known to occur both downslope and in-stream. We report the effects of forest succession on C and N export in the west central Cascades of Oregon, a region of low anthropogenic N input. In a previous study, watersheds with forests of differing ages showed a number of significant differences in stream nutrient export. This study was intended to establish whether differences in stream chemistry were due to variation in N retention by forests of different ages, and thus we measured C and N in lysimeter water draining 12 forest plots, which were categorized into four different stages of successional development. Mean total dissolved nitrogen (TDN) concentrations in deep soil solutions were 2.5 times higher than stream water TDN observed in the previous study, suggesting that denitrification and/or N uptake occurred in the streams or the riparian zone. Although there was a trend for highest soil solution N concentrations in the second youngest (stem exclusion) stage, this trend was significant only for NH₄-N. We previously found that streamwater NO₃-N concentrations averaged 46% of TDN export and was significantly higher in the young than in the older watersheds, however, soil solution NO₃-N concentration averaged 2% of TDN concentration and did not vary with succession. Although NH₄-N concentrations were very low (~5 μg L^?1) in stream water, NH₄-N in lysimeter samples averaged 35% of TDN. While stream water dissolved organic nitrogen (DON) concentrations averaged 30% of TDN concentrations, soil solution DON concentrations averaged 64% of TDN concentration; neither varied with succession. Even with sharp differences in both forest floor and mineral soil C:N ratios and C contents among plots, no measure of N export from the forest stands was significantly related to forest floor or mineral soil characteristics. This is most likely because forest floor C:N ratios all greatly exceeded the reported low C:N ratios required to allow significant N leakage. Taken together, these results suggest that riparian dynamics, in-stream processing, or perhaps even the presence of near-stream alders significantly alter concentrations of all N species between the soil solution and stream water.     

Mycorrhizal associations of dominant trees influence nitrate leaching responses to N deposition

Temperate forests receive some of the highest rates of nitrogen (N) deposition in the world. While numerous studies have investigated the effects of N enrichment on forests, there is little consensus on why some forests become N saturated while others do not. To investigate this, we used a multi-factor meta-analysis to simultaneously estimate the relative importance of several environmental, experimental, and anthropogenic variables on nitrate (NO₃ ^?) leaching in response to experimental N addition. Given that overstory tree species composition and soil C:N ratio influence forest responses to N, we hypothesized that forests dominated by arbuscular mycorrhizal (AM) trees would respond differently than forests dominated by ectomycorrhizal (ECM) trees in the context of forest susceptibility to NO₃ ^? leaching. We found that mycorrhizal association is an important predictor of NO₃ ^? leaching, and AM-dominated forests leach more NO₃ ^? in response to N deposition than ECM forests. Additionally, we found that the amount of total N added, ambient N deposition rates, and the form of N added influenced the magnitude of the NO₃ ^? leaching response. Given that the mycorrhizal associations of most temperate trees are known, our results suggest that this functional grouping may be useful in identifying forests that are most susceptible to NO₃ ^? leaching.     

Multiyear dual nitrate isotope signatures suggest that N‐saturated subtropical forested catchments can act as robust N sinks

In forests of the humid subtropics of China, chronically elevated nitrogen (N) deposition, predominantly as ammonium (NH₄⁺), causes significant nitrate (NO₃^?) leaching from well‐drained acid forest soils on hill slopes (HS), whereas significant retention of NO₃^? occurs in near‐stream environments (groundwater discharge zones, GDZ). To aid our understanding of N transformations on the catchment level, we studied spatial and temporal variabilities of concentration and natural abundance (δ¹⁵N and δ¹⁸O) of nitrate (NO₃^?) in soil pore water along a hydrological continuum in the N‐saturated Tieshanping (TSP) catchment, southwest China. Our data show that effective removal of atmogenic NH₄⁺ and production of NO₃^? in soils on HS were associated with a significant decrease in δ¹⁵N‐NO₃^?, suggesting efficient nitrification despite low soil pH. The concentration of NO₃^? declined sharply along the hydrological flow path in the GDZ. This decline was associated with a significant increase in both δ¹⁵N and δ¹⁸O of residual NO₃^?, providing evidence that the GDZ acts as an N sink due to denitrification. The observed apparent ¹⁵N enrichment factor (ε) of NO₃^? of about ?5‰ in the GDZ is similar to values previously reported for efficient denitrification in riparian and groundwater systems. Episode studies in the summers of 2009, 2010 and 2013 revealed that the spatial pattern of δ¹⁵N and δ¹⁸O‐NO₃^? in soil water was remarkably similar from year to year. The importance of denitrification as a major N sink was also seen at the catchment scale, as largest δ¹⁵N‐NO₃^? values in stream water were observed at lowest discharge, confirming the importance of the relatively small GDZ for N removal under base flow conditions. This study, explicitly recognizing hydrologically connected landscape elements, reveals an overlooked but robust N sink in N‐saturated, subtropical forests with important implications for regional N budgets.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.     

Wildfire Effects on Soil Gross Nitrogen Transformation Rates in Coniferous Forests of Central Idaho, USA

Forest fires often result in a series of biogeochemical processes that increase soil nitrate (NO₃ ^?) concentrations for several years; however, the dynamic nature of inorganic nitrogen (N) cycling in the plant–microbe–soil complex makes it challenging to determine the direct causes of increased soil NO₃ ^?. We measured gross inorganic N transformation rates in mineral soils 2 years after wildfires in three central Idaho coniferous forests to determine the causes of the elevated soil NO₃ ^?. We also measured key factors that could affect the soil N processes, including temperature during soil incubation in situ, soil water content, pH and carbon (C) availability. We found no significant differences (P = 0.461) in gross nitrification rates between burned and control soils. However, microbial NO₃ ^? uptake rates were significantly lower (P = 0.078) in burned than control soils. The reduced consumption of NO₃ ^? caused slightly elevated NO₃ ^? concentrations in the burned soils. C availability was positively correlated with microbial NO₃ ^? uptake rates. Despite reduced microbial NO₃ ^? uptake capacity in the burned soils, soil microbes were a strong enough N sink to maintain low soil NO₃ ^? concentrations 2 years post fire. Soil NH₄ ⁺ concentrations between the treatments were not significantly different (P = 0.673). However, gross NH₄ ⁺ production and microbial uptake rates in burned soils were significantly lower (P = 0.028 and 0.035, respectively) than in the controls, and these rates were positively correlated with C availability. Our results imply that C availability is an important factor regulating soil N cycling of coniferous forests in the region.     

Large Loss of Dissolved Organic Nitrogen from Nitrogen-Saturated Forests in Subtropical China

Dissolved organic nitrogen (DON) has recently been recognized as an important component of terrestrial N cycling, especially under N-limited conditions; however, the effect of increased atmospheric N deposition on DON production and loss from forest soils remains controversial. Here we report DON and dissolved organic carbon (DOC) losses from forest soils receiving very high long-term ambient atmospheric N deposition with or without additional experimental N inputs, to investigate DON biogeochemistry under N-saturated conditions. We studied an old-growth forest, a young pine forest, and a young mixed pine/broadleaf forest in subtropical southern China. All three forests have previously been shown to have high nitrate (NO₃⁻) leaching losses, with the highest loss found in the old-growth forest. We hypothesized that DON leaching loss would be forest specific and that the strongest response to experimental N input would be in the N-saturated old-growth forest. Our results showed that under ambient deposition (35–50 kg N ha⁻¹ y⁻¹ as throughfall input), DON leaching below the major rooting zone in all three forests was high (6.5–16.9 kg N ha⁻¹ y⁻¹). DON leaching increased 35–162% following 2.5 years of experimental input of 50–150 kg N ha⁻¹ y⁻¹. The fertilizer-driven increase of DON leaching comprised 4–17% of the added N. A concurrent increase in DOC loss was observed only in the pine forest, even though DOC:DON ratios declined in all three forests. Our data showed that DON accounted for 23–38% of total dissolved N in leaching, highlighting that DON could be a significant pathway of N loss from forests moving toward N saturation. The most pronounced N treatment effect on DON fluxes was not found in the old-growth forest that had the highest DON loss under ambient conditions. DON leaching was highly correlated with NO₃⁻ leaching in all three forests. We hypothesize that abiotic incorporation of excess NO₃⁻ (through chemically reactive NO₂⁻) into soil organic matter and the consequent production of N-enriched dissolved organic matter is a major mechanism for the consistent and large DON loss in the N-saturated subtropical forests of southern China. Dr. YT Fang performed research, analyzed data, and wrote the paper; Prof. WX Zhu participated in the initial experimental design, analyzed data, and took part in writing the paper; Prof. P Gundersen conceived the study and took part in writing; Prof. JM Mo and Prof. GY Zhou conceived study; Prof. M Yoh analyzed part of the data and contributed to the development of DON model.     

The influence of catchment characteristics on the seasonality of carbon and nitrogen species concentrations in upland rivers of Northern Scotland

Data from 13 catchments with no arable land in Northern Scotland were used to develop empirical linear regression models of average monthly NO₃ ^? concentrations and average summer and winter concentrations for NH₄ ⁺, dissolved organic N (DON) and dissolved organic carbon (DOC) as a function of catchment characteristics. All catchments displayed a pronounced seasonal NO₃ ^? cycle. Variation in monthly mean NO₃ ^? concentration within and between catchments could be predicted from mean monthly air temperature using separate regression equations for temperatures < and ≥ 5 °C. Soil type, climate and land use influenced NH₄ ⁺ concentrations. In summer, concentrations of NH₄ ⁺ were largest in catchments with extensive areas of brown forest soils, which are less acidic and more base-rich than other upland soils. However, concentrations declined with increasing conifer cover and summer rainfall. In winter, however, % conifer cover had a positive effect, while higher temperature and higher humus iron podzol cover had negative influences. DON concentration decreased with increasing catchment elevation in both summer and winter. Surprisingly, concentrations of DON only displayed a positive relationship with percentage peat cover in the summer. The most important factor controlling DOC concentration was soil type, with a positive relationship being observed between DOC and peat and humus iron podzol coverage. Elevation was also important, but only in the winter when concentrations were negatively correlated with maximum catchment elevation. Overall, multivariate regression equations explained the spatial and seasonal variability in N species concentrations over a range of catchments within Northern Scotland.     

An exploration of how litter controls drainage water DIN, DON and DOC dynamics in freely draining acid grassland soils

Surface and subsurface litter fulfil many functions in the biogeochemical cycling of C and N in terrestrial ecosystems. These were explored using a microcosm study by monitoring dissolved inorganic nitrogen (DIN) (NH₄ ⁺–N?+?NO₃ ^?–N), dissolved organic nitrogen (DON) and dissolved organic carbon (DOC) concentrations and fluxes in drainage water under ambient outdoor temperatures. Subsurface litter remarkably reduced the DIN concentrations in winter, probably by microbial N uptake associated with higher C:N ratio of added litter compared with soil at 10–25?cm depth. Fluxes of DIN were generally dominated by NO₃ ^?–N; but NH₄ ⁺–N strongly dominated DIN fluxes during freeze–thaw events. Appreciable concentrations of NH₄ ⁺–N were observed in the drainage from the acid grassland soils throughout the experiment, indicating NH₄ ⁺–N mobility and export in drainage water especially during freeze–thaw. Litter contributed substantially to DOC and DON production and they were correlated positively (p?<?0.01) for all treatments. DOC and DON concentrations correlated with temperature for the control (p?<?0.01) and surface litter (p?<?0.001) treatments and they were higher in late summer. The subsurface litter treatment, however, moderated the effect of temperature on DOC and DON dynamics. Cumulative N species fluxes confirmed the dominance of litter as the source of DON and DOC in the drainage water. DON constituted 42, 46 and 62% of cumulative TDN flux for control, surface litter and subsurface litter treatments respectively.     

Microbial N turnover processes in three forest soil layers following clear cutting of an N saturated mature spruce stand

Microbial N turnover processes were investigated in three different forest soil layers [organic (O) layer, 0–10 cm depth (M₁), 10–40 cm depth (M₂)] after the clear cutting of a nitrogen (N) saturated spruce stand at the Höglwald Forest (Bavaria, Germany). The aim of the study was to provide detailed insight into soil-layer specific microbial production and the consumption of inorganic N within the main rooting zone. Furthermore, we intended to clarify the relevance of each soil layer investigated in respect of the observed high spatial variation of seepage water nitrate (NO ₃ ^? ) concentration at a depth of 40 cm. The buried bag and the ¹⁵N pool dilution techniques were applied to determine the net and gross N turnover rates. In addition, soil pH, C:N ratio, pool sizes of soil ammonium (NH ₄ ⁺ ) and NO ₃ ^? , as well as quantities of microbial biomass carbon (C_mic) and nitrogen (N_mic) were determined. The 40 cm thick upper mineral soil was found to be the main place of NO ₃ ^? production with a NO ₃ ^? supply or net nitrification three times higher than in the considerably thinner O layer. Nevertheless, O layer nitrification processes determined via in situ field experiments showed significant correlation with seepage water NO ₃ ^? . An improved correlation noted several months after the cut may result from a transport-induced time shift of NO ₃ ^? with downstream hydrological pathways. In contrast, the soil laboratory incubation experiments found no indication that mineral soil is relevant for the spatial heterogeneity of seepage water NO ₃ ^? . The results from our study imply that in situ experiments may be better suited to studies investigating N turnover in relation to NO ₃ ^? loss via seepage water in similar ecosystems in order to gain representative data.     

Amino acid uptake in deciduous and coniferous taiga ecosystems

We measured in situ uptake of amino acids and ammonium across deciduous and coniferous taiga forest ecosystems in interior Alaska to examine the idea that late successional (coniferous) forests rely more heavily on dissolved organic nitrogen (DON), than do early successional (deciduous) ecosystems. We traced ¹⁵N-NH₄⁺ and ¹³C-¹⁵N-amino acids from the soil solution into plant roots and soil pools over a 24 h period in stands of early successional willow and late successional black spruce. Late successional soils have much higher concentrations of amino acid in soil solution and a greater ratio of DON to dissolved inorganic N (DIN) (ammonium plus nitrate) than do early successional soils. Moreover, late successional coniferous forests exhibit higher rates of soil proteolytic activity, but lower rates of inorganic N turnover. Differences in ammonium and amino acid uptake by early successional willow stands were insignificant. By contrast, the in situ uptake of amino acid by late successional black spruce forests were approximately 4-fold greater than ammonium uptake. The relative difference in uptake of ammonium and amino acids in these forests was approximately proportional to the relative difference of these N forms in the soil solution. Thus, we suggest that differences in uptake of different N forms across succession in these boreal forests largely reflect edaphic variation in available soil N (composition), rather than any apparent physiological specialization to absorb particular forms of N. These finding are relevant to our understanding of how taiga ecosystems may respond to increases in temperature, fire frequency, N deposition, and other potential consequences of global change.     

Soil Solution Nitrogen Losses During Clearing of Lowland Amazon Forest for Pasture

Losses of nitrogen (N) often follow severe disturbance of forest ecosystems. In tropical forests, losses of N associated with the disturbance of clearing may be particularly important because rates of soil N cycling are high and forest clearing now occurs on a large scale. We measured soil solution inorganic N concentrations and fluxes for 1 year in an intact forest in the Brazilian Amazon state of Rondônia and in an adjacent 3-ha forest plot that was cleared for pasture by cutting, burning and planting pasture grass and in established cattle pastures on the same soils that were 5 and 22 years old. The cleared forest had higher soil solution NO ₃ ^? concentrations than the intact forest, but the difference between the cleared and control forests declined with time after the start of the first post-clearing rainy season. Established pastures had much lower solution NH ₄ ⁺ and NO ₃ ^? concentrations than forest or cleared forest. Estimated annual dissolved inorganic solution N fluxes to below 1 m during the first year after clearing were 2.5 kg ha^?1 in forest and 24.4 kg ha^?1 in newly cleared forest compared with only 0.5–1.2 kg ha^?1 in established pastures. The solution fluxes from cleared forest during the first year after clearing were approximately 7 times greater than gaseous N oxide (N₂O+NO) losses estimated for the same time. These results were consistent with the characterization of moist tropical forests on weathered soils as N-rich and likely to respond to disturbances that elevate soil N availability with increased loss to both soil solution and the atmosphere. These results also suggest that the relative increase in N oxide loss is substantially less than the increase solution inorganic N loss.     

Inorganic nitrogen immobilization in live and sterile soil of old-growth conifer and hardwood forests: implications for ecosystem nitrogen retention

Rapid immobilization of inorganic nitrogen (N) in soil contributes to ecosystem N accumulation, even in old-growth and chronically-fertilized forests once thought to have poor N retention capacity. In old-growth conifer and hardwood stands in Pennsylvania, we tested the hypotheses that biotic and abiotic N immobilization are regulated by N form and forest type. We added ¹⁵NH₄ ⁺, ¹⁵NO₂ ^?, and ¹⁵NO₃ ^? to sterile (γ-irradiated) and live organic-horizon soil and define N immobilization as the mass of added ¹⁵N remaining in soil following extractions conducted 15 min, 24 h, and 21 days later. Immobilization of NO₂ ^? (19–25% of added N) occurred in sterile soils within 15 min and was little changed thereafter. Tracer NO₃ ^? immobilization was not observed, although soils had been pretreated (refrigerated) so as to quantify the lower limit of immobilization potential. Immobilization of NH₄ ⁺ (27%) occurred in live conifer soils by 21 days but not in other treatments. In 21-day incubations, tracer N immobilization was greater in NO₃ ^?-poor and humic-rich soils. Immobilization was greater in sterile than in live soil, perhaps owing to artifacts of sterilization. Conifer stands exhibited more massive O-horizons, so NO₂ ^? immobilization per unit area was greater in conifer (1.46 mg N m^?2) than hardwood (0.43 mg N m^?2) stands, possibly accounting for lower N leaching from conifer forests. Areal immobilization rates appear to be fast enough to retain all N transformed to NO₂ ^?, so NO₂ ^? production may be a limiting step in soil N retention in old-growth ecosystems.     

Controls over Soil Nitrogen Pools in a Semiarid Grassland Under Elevated CO2 and Warming

Long-term responses of terrestrial ecosystems to the combined effects of warming and elevated CO₂ (eCO₂) will likely be regulated by N availability. The stock of soil N determines availability for organisms, but also influences loss to the atmosphere or groundwater. eCO₂ and warming can elicit changes in soil N via direct effects on microbial and plant activity, or indirectly, via soil moisture. Detangling the interplay of direct- and moisture-mediated impacts on soil N and the role of organisms in controlling soil N will improve predictions of ecosystem-level responses. We followed individual soil N pools over two growing seasons in a semiarid temperate grassland, at the Prairie Heating and CO₂ Enrichment experiment. We evaluated relationships of N pools with environmental factors and explored the role of plants by assessing plant biomass, plant N, and plant inputs to soil. We also assessed N forms in plots with and without vegetation to remove plant-mediated effects. Our study demonstrated that the effects of warming and eCO₂ are highly dependent on individual N form and on year. In this water-constrained grassland, eCO₂, warming and their combination appear to impact soil N pools through a complex combination of direct- and moisture-mediated effects. eCO₂ decreased NO₃ ^? but had neutral to positive effects on NH₄ ⁺ and dissolved organic N (DON), particularly in a wet year. Warming increased NO₃ ^? availability due to a combination of indirect drying and direct temperature-driven effects. Warming also increased DON only in vegetated plots, suggesting plant mediation. Our results suggest that impacts of combined eCO₂ and warming are not always equivalent for plant and soil pools; although warming can help offset the decrease in NO₃ ^? availability for plants under eCO₂, the NO₃ ^? pool in soil is mainly driven by the negative effects of eCO₂.